Zoological Studies 44(3): 299-328 (2005)
Review Article
A Review of Mimicry in Marine Fishes
John E. Randall
Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA. E-mail: jackr@hawaii.rr.com
This paper was presented at the 7th Indo-Pacific Fish Conference held in Taipei, Taiwan (May 16, 2005).
(Accepted April 3, 2005)
John E. Randall (2005) A review of mimicry in marine fishes. Zoological Studies 44(3): 299-328. The terms
protective resemblance, Batesian mimicry, Müllerian mimicry, aggressive mimicry, and social mimicry are
defined. Color illustrations are given for 20 examples of protective resemblance in marine fishes. Ninety-eight
cases of mimicry in marine fishes are discussed, and 104 color illustrations are presented in support of 56 of
these. The explanation for the mimicry of the juvenile surgeonfish Acanthurus pyroferus by the angelfish
Centropyge vrolikii, based on different food habits of the mimic, model, and the territorial damselfish
Plectroglyphidodon lacrymatus, is questioned. http://www.sinica.edu.tw/Journals/44.3/299.pdf
Key words: Protective resemblance, Mimicry, Marine fishes.
W
munity for its removal of ectoparasites. It serves
as the model for the mimicking blenny, enabling
the latter to get close enough to nip the fins of reef
fishes.
However, there is still some confusion with
respect to these 2 terms, as noted by Vane-Wright
(1980), who used crypsis for protective resemblance. He explained that it is“probably impossible to provide a“perfect”definition (i.e., mutually
exclusive) for either term.” For the purpose of
this report, protective resemblance is used for an
animal that closely resembles a part of a substratum, a plant, or a sedentary animal such as a
sponge or soft coral. Mimicry applies only to animals that resemble active animals. In addition to
resemblance in color and morphology, the mimic
may adopt a pattern of behavior to enhance the
deception.
Examples of protective resemblance among
fishes are legion. We immediately think of flatfishes that match the surrounding substratum and
quickly change their pattern as they move over a
hen the subject of mimicry is raised, the
first examples that come to mind are in the insect
world. However, a surprising number of mimics
have been discovered among marine fishes, the
subject of this pictorial review.
There is a need to provide a distinction
between protective resemblance and mimicry.
Poulton (1898, quoted from Cott, 1957: 397)
defined them as follows:“In the former an animal
resembles some object which is of no interest to
its enemy, and in so doing is concealed; in the latter an animal resembles an object which is well
known and avoided by its enemy, and in so doing
becomes conspicuous.”From observations of the
Cleaner Wrasse Labroides dimidiatus and the
Mimic Blenny Aspidontus taeniatus, Randall and
,
Randall (1960 : 445) revised Poulton s definition of
mimicry to”... an animal resembles an object
which is well known and is avoided or not preyed
upon by its enemy, and in so doing becomes conspicuous.”Far from being avoided, the Cleaner
Wrasse is sought by other fishes of the reef com-
* To whom correspondence and reprint requests should be addressed. E-mail: jackr@hawaii.rr.com
299
300
Zoological Studies 44(3): 299-328 (2005)
different colored area or the many frogfishes and
scorpionfishes that are wonderfully camouflaged.
In addition to protective coloration, these fishes
may have fleshy cutaneous flaps and tentacles
that augment their concealment. Among the bestknown masters of camouflage is the S a r g a s sumfish Histrio histrio, typically found in floating
masses of Sargassum in the open sea, hence far
more difficult to detect than one photographed in
the Waikiki Aquarium (Fig. 1). Another frogfish,
Antennarius commerson in the Hawaiian Is. (Figs.
2, 3), was first overlooked as just another small
Fig. 1. Histrio histrio, Hawaiian Is. (aquarium photo).
Fig. 2. Antennarius commerson and Porites lobata, Hawai'i.
Fig. 3. Same fish as in Fig. 2, but lifted off the bottom.
Fig. 4. Antennarius commerson, Papua New Guinea.
Fig. 5. Antennarius commerson, Komodo, Indonesia.
Fig. 6. Antennarius commerson, juveniles, Hawaiian Is.
Fig. 7. Antennarius commerson, Hawaiian Is.
Randall -- Mimicry in Marine Fishes
301
coral head (Porites lobata). Photographs of the
same species (Figs. 4-7) depict the range of color
it can exhibit. Frogfishes cannot change their color
pattern quickly, so they may be found over a substratum unlike that of their color pattern. One in
the Waikiki Aquarium in Honolulu changed from
orange to dark reddish brown in about 2 weeks
after being moved to a different tank.
Breder (1946) reviewed those species of fishes that bear a deceptive resemblance to plants.
Randall and Randall (1960) reported 12 examples
of fishes that resemble plant material, among them
the juvenile labrid Novaculichthys taeniourus (Fig.
8), which looks like drifting algae (the deception
enhanced by its seemingly disoriented movement);
Iniistius pavo (Fig. 9), which behaves like a dead
leaf on the bottom, moving with the surge; and
juvenile Platax orbicularis (Fig. 10), which resembles a drifting leaf, often near the surface (the
author has observed the same behavior in the
Tripletail Lobotes surinamensis). Other protectively colored fishes include the seahorse Hippocampus bargibanti (Fig. 11), which matches the
seafan (Muricella sp.) on which it lives; the
astounding Leafy Seadragon Phycodurus eques of
South Australia (Fig. 12); another pipefish
Syngnathoides biaculeatus (Fig. 13), the wrasse
Novaculoides macrolepidotus (Fig. 14), and the
filefish Acreichthys tomentosus (Fig. 15), all of
which blend with seagrass; the ghost pipefish
Solenostomus cyanopterus (Fig. 16), which looks
like a drifting piece of seagrass or like the green
alga Caulerpa taxifolia (Fig. 17); S. paradoxus
(Fig. 18), which resembles the soft coral
Dendronephthya sp. in which it is hiding or in a
crinoid of the genus Comanthus (Fig. 19); and an
undescribed clingfish of the genus Discotrema
(Fig. 20), which is nestled in its host crinoid.
The best-known category of mimicry is
Batesian. It was discovered by Henry Walter
Bates, who at the age of 23 y explored the
Brazilian forest and observed some butterflies of 2
different families that were remarkably alike. The
common heliconiine butterflies, although fragile
and slow flying, exude noxious secretions and are
not preyed upon by birds. Resembling them are
the less-common pierids that lack such repellent
substances. Bates read his paper to the Linnean
Society in London in 1861.
Müllerian mimicry is named for Fritz Müller
who also studied butterflies in Brazil. Bates found
examples of pairs of inedible butterflies that were
Fig. 8. Novaculichthys taeniourus, juvenile, Batu Ata, Indonesia.
Fig. 9. Iniistius pavo, juvenile, Hawaiian Is.
Fig. 10. Platax orbicularis, juvenile, Sulawesi, Indonesia.
Fig. 11. Hippocampus bargibanti, Sulawesi, Indonesia.
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Zoological Studies 44(3): 299-328 (2005)
not related but looked much alike. Müller (1878)
stated that species with inedible properties may
evolve to look alike. Fewer individuals of each are
sacrificed before predators learn to avoid them
,
(although a predator s aversion may also be
innate).
Cott (1957: 398) differentiated Batesian and
Müllerian mimicry as follows:“In Batesian mimicry
a relatively scarce, palatable, and unprotected
species resembles an abundant, relatively unpalatable, or well-protected species, and so becomes
disguised. In Müllerian mimicry, on the other hand,
a number of different species, all possessing
aposematic attributes and appearance, resemble
one another, and so become more easily recognized.”
The concept that Batesian mimics must be
less abundant than their models has been repeated so frequently in the literature that it would seem
obligatory. However, Fisher (1930) stated that a
Batesian mimic may be more numerous if the
model is extremely noxious or if the mimic is a relatively unimportant item of prey. Springer and
Smith-Vaniz (1972) discussed 7 possible cases
where the mimic may be more abundant. Two of
these were shown experimentally to apply when
the model was a species of Meiacanthus, all of
which are believed to have a venomous bite.
Longley (1917) warned that the similarity of 1
fish to an unrelated species does not necessarily
mean that mimicry is involved. Mere resemblance
might be the case of the cardinalfish Fowleria sp.,
suggested as being a mimic of scorpionfishes by
Siegel and Adamson (1983), or the similar example of Fowleria abocellata (= F. vaiuli) advocated
as mimicking Scorpaenodes guamensis by Goren
and Karplus (1983). More convincing is the apparent mimicry of a scorpionfish by the percoid fish
Centrogenys vaigiensis, as discussed by Whitley
(1935). This fish was first described by Quoy and
Gaimard (1824) as a species of Scorpaena.
Cuvier in Cuvier and Valenciennes (1829) realized
it was not a scorpionfish and renamed it
Centropristes scorpenoides, but Quoy and
,
Gaimard s species name prevails. The similarity
may be seen in figure 21 of C. vaigiensis and fig-
Fig. 13. Syngnathoides biaculeatus, Papua New Guinea.
Fig. 14. Novaculoides macrolepidotus, Cebu, Philippines.
Fig. 12. Phycodurus eques, South Australia (aquarium photo).
Fig. 15. Acreichthys tomentosus, Papua New Guinea.
Randall -- Mimicry in Marine Fishes
303
Figs. 16-17. Solenostomus cyanopterus, Mai I., Indonesia
(left); Sulawesi, Indonesia (right; John L. Earle).
Figs. 18-19. Solenostomus paradoxus, Alor, Indonesia (left);
Papua New Guinea (right).
Fig. 20. Discotrema sp., New Britain.
Fig. 21. Centrogenys vaigiensis, Sulawesi, Indonesia.
Fig. 22. Scorpaenodes guamensis, Mauritius.
Fig. 23. Myrichthys colubrinus, Bali, Indonesia.
Fig. 24. Myrichthys colubrinus, Tuamotu Archipelago (Yves
Lefevre).
Fig. 25. Laticauda colubrina, Banda Sea (Ron and Valerie
Taylor).
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Zoological Studies 44(3): 299-328 (2005)
ure 22 of the scorpionfish Scorpaenodes guamensis.
An early example of Batesian mimicry in fishes is the resemblance of the harmless Ringed
Snake Eel Myrichthys colubrinus (Figs. 23, 24) to
the venomous sea snake Laticauda colubrina (Fig.
25) and other dark-ringed sea snakes (discussed
by McCosker and Rosenblatt 1993). Unlike other
Indo-Pacific snake eels, in general, this one roams
freely in the open by day. The width of the dark
bars on the snake eel varies, as shown by the
extreme examples of figures 23 and 24. A photograph of a snake eel was recently sent to the
author from Tahiti to determine if it was a sea
snake. It proved to be another snake eel,
Leiuranus semicinctus, a less-likely mimic of a sea
snake because the dark bars are broader than the
pale interspaces and do not completely encircle
the body.
Mahadevan and Nagappan Nayar (1965)
observed the yellow and black-barred juvenile
carangid fish Gnathanodon speciosus closely
associated with a sea snake of the same color pattern in southeastern India. When approached, the
fish“moved closer to the body of the snake, literally hugging the body.”They added that it was difficult to see the fish at first sight because its color
pattern blended with that of the snake. It should
be noted, however, that juveniles of G. speciosus
often swim at the flank of large bony fishes or ride
the bow wave of sharks.
Masterman (1908) reported the similarity of
the black-marked pectoral fin of the Common Sole
Solea vulgaris in Europe to the black 1st dorsal fin
of the venomous weeverfishes Echiichthys vipera
and Trachinus draco.
The juvenile of the carangid fish Alectis ciliaris
(Fig. 26) has long been presumed to be a mimic of
venomous jellyfishes. The very long anterior soft
dorsal and anal rays of Alectis resemble the long
thread-like tentacles of virulent cubomedusae. It is
not known to associate closely with jellyfishes as
do juveniles of some other carangid fishes, stromateoids, etc. (Mansueti 1963).
Schooling juveniles of the Convictfish
Pholidichthys leucotaenia (Fig. 27) look very much
like schools of the juvenile venomous catfish
Plotosus lineatus (Fig. 28). The catfish aggregate
closely for the mutual protection provided by their
spines, and Convictfish juveniles may also form
dense schools, although they tend to be more dispersed.
Longley and Hildebrand (1940) described
Hemiemblemaria simulus as a new genus and
species of the pike blenny family Chaenopsidae
from Tortugas, Florida. They regarded the
chaenopsid as a Batesian mimic of the wrasse
Thalassoma bifasciatum. It was observed to leave
the bottom and swim freely with the model, using
its pectoral fins like a labrid. Thalassoma bifasciatum is well known to feed in part on the ectoparasites of other fishes, therefore enjoying protection
from predation (though not complete, because a
few have been found in the stomachs of predaceous fishes). This behavior provides the opportunity for the chaenopsid to safely feed on zooplank-
Fig. 26. Alectis ciliaris, juveniles, Hawaiian Is.
Fig. 27. Pholidichthys leucotaenia, Papua New Guinea.
Fig. 28. Plotosus lineatus, Cebu, the Philippines.
Randall -- Mimicry in Marine Fishes
305
ton as it mingles with the labrid. Randall and
Randall (1960) reported on the food habits of the
chaenopsid (free-living copepods, mysids, and a
few small fishes) from specimens collected in the
Florida Keys, and they illustrated a juvenile and an
adult of both the mimic and the model (the initialphase adult of the model, not the larger and very
differently colored terminal male).
Randall and Emery (1971) were completely
deceived by a juvenile of the ephippid fish Platax
pinnatus (Fig. 29) that they observed and collected
in Palau in 1970. They thought it was a turbellarian flatworm because of its coloration and behavior.
When on the bottom, its locomotion was like that of
a flatworm, and when above the bottom, it oriented
horizontally and swam slowly with an undulating
motion. Several polyclad flatworms are dark with
an orange margin or submarginal band. An example is given here of a possible model, Pseudoceros
periaurantus (Fig. 30). Some polyclad flatworms
are known to be strongly toxic and advertise this
with bright warning coloration. Fishes either avoid
them, or if they ingest them, they quickly spit them
out. Exceptions are puffers, many of which are
well known to be toxic themselves (Newman and
Cannon 2003).
A small sole (Fig. 31) from Papua New
Guinea, estimated to be 12 mm in length by the
photographer, appears to be a juvenile of an undetermined species of Soleichthys. It moves slowly
over the bottom like a flatworm and is believed to
be a mimic of an undescribed polyclad of the
genus Pseudoceros (Fig. 32). Kuiter (1991) first
illustrated this juvenile sole and noted its movement like a flatworm. Newman and Cannon (2003)
also figured it, as well as its polyclad model.
Heck and Weinstein (1978) observed the
striking resemblance of the juvenile of the burrfish
Chilomycterus antennatus to the sea hare Aplysia
dactylomedia in seagrass meadows of the
Caribbean Sea. They concluded that the burrfish
is a Batesian mimic of the unpalatable sea hare.
Kuiter (1991: 116) illustrated“a small anglerfish clinging to rocks like a nudibranch which as a
predator can surprise prey and has little fear of
being preyed upon as nudibranchs are reported to
be of bad taste.” His photograph appears to be a
juvenile of the frogfish Antennarius pictus.
Remarkable examples of Batesian mimicry
are those blennies of the genera Ecsenius,
Petroscirtes, and Plagiotremus that mimic fangblennies of the genus Meiacanthus (Springer and
Fig. 29. Platax pinnatus, Sulawesi, Indonesia (Mike Severns).
Fig. 30. Pseudoceros periaurantias (Leslie Newman and A.
Flowers).
Fig. 31. Soleichthys sp., Papua New Guinea (David Hall).
Fig. 32. Pseudoceros sp., Guam (Gustav Paulay).
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Zoological Studies 44(3): 299-328 (2005)
Smith-Vaniz 1972, Losey 1972, Russell 1975,
Allen et al. 1975, Smith-Vaniz 1976 1987, Russell
et al. 1976, Smith-Vaniz et al. 2001). Species of
Meiacanthus have venomous glands associated
with the pair of very large grooved canine teeth in
the lower jaw. Species of other genera of fangblennies, such as Aspidontus, Petroscirtes, and
Plagiotremus, also have a pair of large canines in
the lower jaw, but they are not venomous. The
canines are used in defense and infraspecific
aggression, not for feeding (Russell 1977: fig. 2).
Springer and Smith-Vaniz (1972) performed
experiments by feeding the Red Sea blennies
Meiacanthus nigrolineatus, Ecsenius gravieri, and
Plagiotremus townsendi to scorpionfishes and
groupers in laboratory tanks in the Gulf of Aqaba.
They concluded that“live Meiacanthus nigrolineatus is a generally unacceptable prey species,
whose unacceptability lies in its bite.” Ecsenius
gravieri and P. townsendi are generally acceptable
prey species, but they are often not eaten when
the predator has previously ingested and rejected
M. nigrolineatus.
Three cardinalfishes of the genus Cheilodipterus and juveniles of the breams Scolopsis
bilineatus and S. margaritifera are also mimics of
species of Meiacanthus. Russell (1975) wrote of
the Scolopsis,“Although deeper bodied than the
blenny, the belly is white, in pale contrast to the
colours above, so that at first sight, especially
when viewed from above, the small coral bream
appears to be a blenny.” He pointed out that the
mimicry is further enhanced by the juvenile
,
bream s swimming in the same distinctive way as
species of Meiacanthus. The blennies swim a
short straight distance, pause, then make another
linear swim, often in a different direction. The
mimicking cardinalfishes of the genus Cheilodipterus swim the same way. This is remarkable
because species of the family Apogonidae are
generally nocturnal. The bream mimicry is most
convincing because juveniles of the same species
exhibit a different color pattern to correspond to
that of species of Meiacanthus in the area. The
following figures show model species of
Meiacanthus and their mimics.
In figure 33, Meiacanthus nigrolineatus and its
mimic, Ecsenius gravieri, were photographed
together in the Gulf of Aqaba, Red Sea. Meiacanthus nigrolineatus is darker in the southern
part of the Red Sea; E. gravieri closely matches its
color there. Unlike other species of Ecsenius,
Fig. 33. Meiacanthus nigrolineatus (left) and Ecsenius gravieri,
Red Sea (Victor G. Springer).
Fig. 34. Plagiotremus townsendi (Robert F. Myers).
Fig. 35. Meiacanthus atrodorsalis, Banda, Indonesia.
Fig. 36. Plagiotremus laudandus, Wetar, Indonesia.
Randall -- Mimicry in Marine Fishes
307
which rest upon the substratum except when moving to a new location or feeding, E. gravieri swims
freely above the bottom like M. nigrolineatus.
Figure 34 of Plagiotremus townsendi in the
Gulf of Aqaba also shows that it mimics M. nigrolineatus. To offset its more-slender body, it tends
to hold its dorsal and anal fins erect. The same is
true of other mimic species of Plagiotremus.
In figure 35, Meiacanthus atrodorsalis, wideranging in the Pacific from Japan to Australia and
east to the islands of Samoa, is mimicked by
Plagiotremus laudandus (Fig. 36). The bluish
Fig. 37. Meiacanthus oualanensis, Fiji.
Fig. 38. Plagiotremus laudandus flavus, Fiji.
Fig. 39. Scolopsis bilineatus, juvenile, Fiji.
Fig. 40. Scolopsis bilineatus, adult, Luzon, the Philippines.
Fig. 41. Meiacanthus smithi, Maldive Is.
Fig. 42. Plagiotremus phenax, Sumatra, Indonesia.
Fig. 43. Scolopsis bilineatus, juvenile, Sumatra, Indonesia.
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Zoological Studies 44(3): 299-328 (2005)
brown and orange color phase of Ecsenius bicolor
has been regarded by some authors as a mimic of
M. atrodorsalis, but it is a marginal example at best
(Springer 1988).
The yellow Meiacanthus oualanensis (Fig. 37)
found in Fiji serves as the model for the yellow
Plagiotremus laudandus flavus (Fig. 38) (William F.
Smith-Vaniz, pers. comm., now prefers to regard
flavus as a species) and the yellow-phase juvenile
of the Bridled Bream Scolopsis bilineatus (Fig. 39).
Four different color forms of the bream mimic different species of Meiacanthus, but the color of the
adult bream (Fig. 40) is essentially the same
throughout its broad range.
In figure 41, Meiacanthus smithi in the Indian
Ocean is mimicked by Plagiotremus phenax (Fig.
42), and a different-colored juvenile of Scolopsis
bilineatus (Fig. 43). The species name, phenax, is
from the Greek for imposter, hence well chosen for
the mimicking Plagiotremus.
Meiacanthus grammistes from southern
Japan to Australia and the Solomon Is. (Fig. 44) is
the model for the fangblenny Petroscirtes breviceps (Fig. 45), the cardinalfish Cheilodipterus
nigrotaeniatus (Fig. 46), and a third color form of
juvenile Scolopsis bilineatus (Fig. 47).
Smith-Vaniz et al. (2001: fig. 15) illustrated a
similar mimic-model group, Meiacanthus lineatus,
Petroscirtes fallax, and juvenile Scolopsis bilineatus, but their illustration of P. fallax shows 2 white
instead of bright yellow stripes. However, SmithVaniz (pers. comm.) has seen photographs of P.
fallax in which the stripes are bright yellow).
Smith-Vaniz et al. cited Yatsu et al. (1983), who
mentioned the similarity of Petroscirtes breviceps
to Meiacanthus kamoharai as a possible mimetic
pair. Smith-Vaniz et al. also suggested that the
young of the nemipterid fish Pentapodus trivittatus
is a mimic of Meiacanthus crinitus. Lacking a photograph of the juvenile Pentapodus, they presented one of a subadult, and the resemblance is
clear.
Meiacanthus geminatus from Sabah and the
Philippines (Fig. 48) is the model for Cheilodipterus zonatus (Fig. 49) and the young of
Scolopsis margaritifera (Fig. 50). An adult of
Scolopsis is shown in figure 51.
Meiacanthus vittatus from Papua New Guinea
(Fig. 52) is mimicked by Petroscirtes breviceps
(Fig. 53; note the difference from P. breviceps of
Fig. 45), by Cheilodipterus parazonatus (Fig. 54),
and by another color form of the juvenile of
Fig. 44. Meiacanthus grammistes, Alor, Indonesia.
Fig. 45. Petroscirtes breviceps, Sulawesi, Indonesia.
Fig. 46. Cheilodipterus nigrotaeniatus, Sulawesi, Indonesia
(Gerald R. Allen).
Fig. 47. Scolopsis bilineatus, juvenile, Luzon, the Philippines.
Randall -- Mimicry in Marine Fishes
309
Fig. 48. Meiacanthus geminatus, Papua New Guinea (Gerald
R. Allen).
Fig. 49. Cheilodipterus zonatus, Papua New Guinea (Gerald
R. Allen).
Fig. 50. Scolopsis margaritifera, juvenile, Palau.
Fig. 51. Scolopsis margaritifera, adult, Sulawesi, Indonesia.
Fig. 52. Meiacanthus vittatus, Papua New Guinea.
Fig. 53. Petroscirtes breviceps, Papua New Guinea.
Fig. 54. Cheilodipterus parazonatus, Papua New Guinea.
Fig. 55. Scolopsis margaritifera, Solomon Is. (Gerald R. Allen).
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Zoological Studies 44(3): 299-328 (2005)
Fig. 56. Meiacanthus anema, Ambon, Indonesia.
Fig. 57. Amblygobius linki, Ambon, Indonesia.
Scolopsis margaritifera (Fig. 55).
Springer and Smith-Vaniz (1972) found a
museum jar containing a mixture of Meiacanthus
anema (Bleeker) and the goby Amblygobius linki
Herre from Buru, Indonesia that had been sorted
as 1 species. Both the blenny and the goby have
a similar black-striped color pattern in preservative.
They wrote,“these two similar species could possibly be involved in a mimetic relationship.”The
venomous fangblenny would therefore be the
model for the mimicking goby. However, they
added that the color illustration of M. anema by
Bleeker from the type locality of Ambon, Indonesia
showed the pale spaces between the black stripes
as yellow, whereas Herre described the color of A.
linki from the type locality of Bungau, Sulu
Province, Philippines as blue with black stripes.
The author collected both species from a brackish
area in Ambon Bay, and his photographs (shown
here as Figs. 56 and 57) support the suggestion of
Batesian mimicry. If Amblygobius linki is blue and
black in the Philippines, it would be of interest to
know if Meiacanthus anema occurs in the same
area and has the same color. Also it would be
more convincing as an example of mimicry if the
goby were known to adopt a mode of swimming
like the blenny.
Sazima (2002a) presented convincing evidence with color illustrations to show that the juvenile of the haemulid Pomadasys ramosus in southeastern Brazil mimics the juvenile of the carangid
Oligoplites palometa, a species with venom glands
in its dorsal and anal spines. He also summarized
24 examples of Batesian mimicry in marine fishes
(his table 1).
Springer and Smith-Vaniz (1972) pointed out
that the model-mimic pairs of species of
Meiacanthus and Plagiotremus are also Müllerian
mimics. Fishes do not prey upon species of
Meiacanthus because of the venomous bite they
can expect, and they try to prevent the species of
Plagiotremus from approaching within striking
range. Both species of fangblennies of a mimicmodel pair are avoided by predators; therefore
their protection is enhanced by their similarity.
Aggressive mimicry (Wickler 1965 1968),
sometimes known as Peckhamian mimicry (after
E.G. Peckham, who discovered it while studying
spiders), is used to depict an animal or a part
thereof that enables a predator to get closer to its
prey or to attract its prey to within striking range.
Therefore, the Batesian blenny mimic Aspidontus
taeniatus is also an aggressive mimic (Batesian in
resembling the protected Cleaner Wrasse
Labroides dimidiatus, and aggressive when it bites
pieces from the fins of other fishes). The same is
true of the mimic species of fangblennies (also
called sabertooth blennies) of the genus
Plagiotremus. Most of the other aggressive mimics operate as mimics only in the juvenile phase.
They change to the different adult coloration when
they become larger than their models.
Hobson (1969) reported that Plagiotremus
azaleus, the only fangblenny from the eastern
Pacific, aggregates with the labrid Thalassoma
lucasanum. Although not closely resembling the
wrasse, this deceptive behavior improves the success of its attacks on reef fishes for mucus and
epidermal tissue. Russell et al. (1976) observed
Plagiotremus tapeinosoma schooling with the
labrid fish Thalassoma amblycephalum, the plesiopid Trachinops taeniatus (Fig. 58), and the
tripterygiid Forsterygion sp. for the same advantage. They also reported individuals of an unusual
orange color morph of Plagiotremus rhinorhynchos
swimming with an aggregation of orange Anthias
mortoni (now Pseudanthias huchtii) and actively
feeding on zooplankton in the passing current.
Randall -- Mimicry in Marine Fishes
311
Fig. 58. Plagiotremus tapeinosoma in school of Trachinops
taeniatus, New South Wales (Barry C. Russell).
Fig. 59. Iracundus signifer, Hawaiian Is.
They wrote:“From amongst the group of anthiines
P. rhynorhynchos launched a series of attacks
against passing fishes, ...”William F. Smith-Vaniz
(pers. comm.) made a similar observation in Fiji.
While watching a feeding aggregation of yellow
Pseudanthias squamipinnis, with about 6 Ecsenius
midas swimming with them (discussed below as
an example of social mimicry), he was surprised to
see 2 Plagiotremus rhinorhynchos in the group, 1
entirely yellow and the other with only a trace of
the usual blue stripes.
Perhaps the best-known examples of aggressive mimicry are found in the frogfish family
Antennariidae (and other ceratioids) that use a
modified 1st dorsal ray as a lure called the illicium.
The esca (bait) at the tip of the illicium can look
very much like a small fish, worm, shrimp, or octopus and serves to attract prey to within striking distance of the mouth of the frogfish (Pietsch and
Grobecker 1978 1987). Similarly, the slender lure
from tissue inside the lower jaw of some of the
stargazers (Uranoscopus) is enticingly wriggled
like a worm from the mouth of an otherwise hidden
stargazer. One snake eel, Glenoglossa wassi
McCosker, 1982, has adopted a comparable oral
lure. Also, the elongate frilled 1st dorsal ray of the
flatfish Asterorhombus fijiensis has been shown to
serve as a lure (Amaoka et al. 1994), as has that
of A. intermedius, reported by Manabe and
Shinomiya (1998).
An unusual example of luring behavior was
discovered in the Indo-Pacific scorpionfish,
Iracundus signifer (Fig. 59), by Shallenberger and
Madden (1973). The dorsal fin looks very much
like a small fish. The black spot on the 2nd fin
membrane is the eye (which enlarges during luring), the gap between the 1st and 2nd spines is
the mouth, and the long 4th dorsal spine is the 1st
dorsal spine of the lure. They wrote:“Typically,
the lure was snapped from side to side. The first
two spines of the spinous dorsal initiated this
movement and the rest of the fin followed in a
sinuous wave down the back.” The 1st and 2nd
dorsal spines are alternately closed together then
separated, giving the effect of respiration. The
resemblance to a small moving fish is remarkable
(the authors showed me a motion picture film of
the behavior). The prey included apogonid and
other scorpaenid fishes up to 1/2 the body length
of the predator.
Fig. 60. Antennarius striatus and Astropyga radiata, Mindoro,
the Philippines (Frank Schneidewind).
Schneidewind (2004) provided an example of
the frogfish Antennarius striatus as a Batesian
mimic of the very venomous sea urchin Astropyga
radiata (Fig. 60). Sea urchins of the genus
Diadema would be a closer model in color. The
esca (“bait”) of the frogfish looks like a succulent
white or pink worm, so this frogfish is also an
aggressive mimic. Sazima (2002b), however, preferred not to regard lures from part of the body of
predatory fishes as true examples of aggressive
mimicry.
Three of the western Atlantic butter hamlets
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Zoological Studies 44(3): 299-328 (2005)
Fig. 61. Chromis cyanea, Bahamas.
Fig. 62. Hypoplectrus gemma, Florida, USA (Paul Humann).
Fig. 63. Microspathodon chrysurus, Belize.
Fig. 64. Hypoplectrus chlorurus, Puerto Rico.
Fig. 65. Stegastes diencaeus, Belize.
Fig. 66. Hypoplectrus nigricans, Bahamas.
(Hypoplectrus spp.) are regarded as aggressive
mimics of harmless damselfishes and thereby get
closer to their prey (Randall and Randall 1960).
Chromis cyanea (Fig. 61) is the model for H.
gemma (Fig. 62); Microspathodon chrysurus (Fig.
63) is the model for H. chlorurus (Fig. 64); and uniformly dark brown species of Stegastes such as S.
diencaeus (Fig. 65) are the models for H. nigricans
(Fig. 66). Thresher (1978) added 3 other mimetic
pairs involving Hypoplectus, including the young of
the angelfish Holacanthus tricolor which serves as
the model for Hypoplectrus guttavarius. Domeier
(1994) suggested that these are just chance
resemblances, noting that the hamlets do not
appear to mimic the behavior of any of the models.
However, he added:“It is possible that the recent
evolutionary history of Hypoplectrus has not
allowed for the complete development of typical
mimic characteristics.” Species of Hypoplectrus
feed more on crustaceans than on fishes (Randall
1967). Thresher pointed out that the crustacean
compound eye, while very sensitive to movement
and overall color, does not permit good form
vision. Except for the final attack, hamlets move
Randall -- Mimicry in Marine Fishes
313
Fig. 67. Lutjanus bohar, juvenile, Sulawesi, Indonesia.
Fig. 68. Chromis ternatensis, Sangihe Is., Indonesia.
Fig. 69. Lutjanus bohar, juvenile, Fiji (Gerald R. Allen).
Fig. 70. Chromis margaritifer, Indonesia.
Fig. 71. Lutjanus bohar, large juvenile, Tuamotu Archipelago.
Fig. 72. Chromis iomelas, Tuamotu Archipelago.
slowly toward their prey. A general resemblance in
color and general shape to a harmless damselfish
model would be expected to result in greater success for a predator of crustaceans.
Russell et al. (1976) reported the juvenile of
the piscivorous snapper Lutjanus bohar (Fig. 67) to
be an aggressive mimic of the damselfish Chromis
ternatensis (Fig. 68), both wide-ranging in the
Indo-Pacific. They found a different color form of
L. bohar (Fig. 69) in Samoa, with Chromis margaritifer serving as the model (Fig. 70). Lutjanus
bohar of figure 71 is mimicking still another white-
tailed Chromis, C. iomelas (Fig. 72), from the
South Pacific. The snapper of figure 72 has
reached a size larger than its model, so it is starting to assume the adult color pattern. Moyer
(1977) wrote that juveniles of L. bohar seem to
mimic the following species of Chromis in Japan:
C. flavomaculata, C. weberi, C. lepidolepis, and C.
miyakeensis (a synonym of C. notata).
Russell et al. (1976) were also the first to publish the amazing similarity of the juvenile grouper
Anyperodon leucogrammicus (Fig. 73) to female
wrasses of the genus Halichoeres. Their choice of
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Zoological Studies 44(3): 299-328 (2005)
Fig. 73. Anyperodon leucogrammicus, juvenile, Sulawesi,
Indonesia.
Fig. 74. Halichoeres leucurus, female, Palau.
Fig. 75. Anyperodon leucogrammicus, juvenile, Sri Lanka
(R. Charles Anderson).
Fig. 76. Halichoeres timorensis, female, Sri Lanka.
Fig. 77. Epinephelus multinotatus, juvenile, Kuwait.
Fig. 78. Neopomacentrus sindensis, Bahrain.
Fig. 79. Oxycheilinus mentalis, Red Sea.
Fig. 80. Oxycheilinus mentalis and Parupeneus macronemus,
Red Sea (Richard Field).
Randall -- Mimicry in Marine Fishes
315
Fig. 81. Epibulus insidiator, female, Red Sea.
Fig. 82. Pomacentrus sulfureus, Red Sea.
the model, H. biocellatus, is not as close a match
as the females of H. leucurus (H. purpurescens is
a synonym) (Fig. 74), H. melanurus, H. richmondi,
and H. vrolickii, as shown by Randall and Kuiter
(1989).
The documentation of mimicry is enhanced
when the mimic exhibits a color pattern to resemble a different model. Such is the case of juvenile
Anyperodon leucogrammicus in Sri Lanka (Fig.
75), where it mimics the female of Halichoeres timorensis (Fig. 76), very different in color from
species of the H. leucurus complex.
Randall (1995) proposed that the juvenile of
the grouper Epinephelus multinotatus (Fig. 77) is a
mimic of the damselfish Neopomacentrus sindensis (Fig. 78) of the Persian Gulf and Arabian Gulf,
as first suggested by Nigel Downing (pers.
comm.).
Kuiter (1995) regarded the juvenile of the
grouper Plectropomus oligacanthus as a mimic of
the wrasse Cheilinus celebicus (now classified in
Oxycheilinus) from underwater observation in the
Seribu Is. north of Jakarta. However, his model is
also a predator with“a similar diet.” He noted
that the grouper of the same size as the wrasse
has a larger mouth, hence would take larger prey
than Oxycheilinus. His figure of the Plectropomus
is not very convincing as being similar in color or
shape to Oxycheilinus celebicus. I regard this as a
questionable example of aggressive mimicry.
Oxycheilinus mentalis of the Red Sea and
western Indian Ocean, a close relative of O.
celebicus, is a mimic (not a model for an aggressive mimic), as shown by Ormond (1980), who
misidentified it as Oxycheilinus digrammus. He
wrote that it is“able to rapidly change its color to
match that of the fish close to which it is
swimming”and gave examples of nonpredatory
parrotfishes and surgeonfishes with which it
swims. Figure 79 shows its non-mimetic color pattern. When O. mentalis is close to the goatfish
Parupeneus macronemus (Fig. 80), it quickly
becomes pale with a midlateral dark stripe.
Ormond added,“It appears to feed on small fish,
fish larvae and perhaps crustaceans.”
Ormond provided another example of a labrid
fish as an aggressive mimic, the Indo-Pacific
Slingjaw Wrasse Epibulus insidiator. Resident
herbivorous fishes of inshore Red Sea reefs, such
as Acanthurus sohal, Plectroglyphidodon lacrymatus, and Stegastes nigricans, vigorously defend
the algal substratum of their territory. The surgeonfish Zebrasoma desjardinii forms feeding
aggregations in order to invade this territory.
Ormond wrote,“An adult E. insidiator, on seeing
such a school of Z. veliferum [the Red Sea/western Indian Ocean species of Sailfin Tang is now Z.
desjardinii], will typically swim over to join it, at the
same time changing colouration to an overall dark
brown-black, similar to that of the Zebrasoma.
Within the middle of a Zebrasoma school, Epibulus
is thus difficult to distinguish from the other fish
(see Plate II); its hunting approaches to the substrate appear much the same as the grazing
movements of Zebrasoma. Possibly Epibulus may
even take some of the smaller pomacentrids that
are attempting to defend their territories.”
One female color form of Epibulus insidiator is
entirely bright yellow or yellow with a narrow black
bar on each scale of the body (Fig. 81). It is
regarded as a probable mimic of the yellow damselfish Pomacentrus sulfureus (Fig. 82) of the Red
Sea and western Indian Ocean (Field, 1997).
Perhaps the same strategy is used to approach
other yellow damselfishes of the genus
Pomacentrus, such as P. pikei of the Mascarene
Is., P. moluccensis of the eastern Indian Ocean
and western Pacific, or Stegastes aureus of the
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Zoological Studies 44(3): 299-328 (2005)
Fig. 83. Mycteroperca tigris, juvenile, Anguilla (Paul Humann).
Fig. 84. Thalassoma bifasciatum, initial phase, Virgin Is.
Fig. 85. Aethaloperca rogaa, juvenile, Bali, Indonesia.
Fig. 86. Centropyge fisheri, Papua New Guinea.
South Pacific. Just comparing photos of Epibulus
and damselfishes, one might be skeptical that
mimicry is involved. However, in aggressive mimicry, the mimic does not have to precisely duplicate
the color and shape of the model to gain the
advantage of getting closer to its prey (Roberts
1990). A Batesian mimic generally needs to be a
more-exact replica of its model because the
mimetic role is to avoid predation.
Sikkel and Hardison (1992) concluded from
quantitative observations that the Yellowtail
Snapper Ocyurus chrysurus is an aggressive
mimic of the Yellow Goatfish Mulloidichthys martinicus, both wide-ranging in the tropical and subtropical western Atlantic. Both are primarily white
with yellow fins and a yellow stripe from the eye to
the caudal fin. The snapper, a predator on a variety of fishes and benthic invertebrates (chiefly
crustaceans), has greater foraging success when it
accompanies the goatfish. Not only does it
approach its prey more readily in its guise of the
goatfish, but it feeds on fishes and crustaceans
flushed by the rooting of the goatfish in the substratum.
Snyder (1999) discovered that the juvenile of
the grouper Mycteroperca tigris (Fig. 83) is a mimic
of the juvenile and initial phase of the abundant
Thalassoma bifasciatum (Fig. 84) in coloration,
size, and swimming mode. Both species are widely distributed in the western Atlantic. He wrote,
“By imitating a non-predatory cleaner fish, the
mimic appears to enjoy a twofold advantage - it
both facilitates prey capture and avoids predation.”
Sazima (2002b) found 2 other groupers of the
genus Mycteroperca that are aggressive mimics of
wrasses from his observations in southeastern
Brazil. Halichoeres maculipinna (whose name was
changed to H. penrosei by Rocha 2004 for this
Brazilian sister species) is the model for
Mycteroperca interstitialis, and H. poeyi is the
model for M. acutirostris.
Snyder et al. (2001) reported the aggressive
mimetic relationship of the juvenile of the IndoPacific grouper Aethaloperca rogaa (Fig. 85) with
dark species of angelfishes of the genus
Centropyge, such as C. fisheri (C. flavicauda is a
synonym) (Fig. 86) and dark damselfishes of the
genera Pomacentrus and Stegastes.
Still another grouper mimic is the juvenile of
the Indo-Pacific Plectropomus laevis (Fig. 87),
which closely resembles the small toxic toby
Canthigaster valentini (Fig. 88) (Randall 2005).
Randall -- Mimicry in Marine Fishes
Fig. 87. Plectropomus laevis, juvenile, Ogasawara Is. (Hajime
Masuda).
Fig. 88. Canthigaster valentini, Great Barrier Reef.
Fig. 89. Paraluteres prionurus, Tonga.
Fig. 90. Paraluteres arqat, Red Sea (Richard Field).
Again, there is a dual basis for the mimicry.
Predators do not target species of Canthigaster
and may therefore overlook the small grouper.
Furthermore, by resembling the toby, the grouper
can approach its prey more closely.
The Indo-Pacific filefish Paraluteres prionurus
(Fig. 89) also mimics Canthigaster valentini, as
shown by Clark and Gohar (1953), Tyler (1966),
and Caley and Schluter (2003). The Red Sea
Paraluteres arqat (Fig. 90) is a mimic of
Canthigaster margaritata (Fig. 91), as first suspected by Clark and Gohar (1953). The filefishes of
the genus Paraluteres are usually seen with the
1st dorsal fin fully depressed; once this fin is erect,
the effect of mimicry is reduced. The 2nd dorsal
fin of Paraluteres is broader than that of
Canthigaster, but it is nearly transparent. Males of
Paraluteres have 2 pairs of spines on each side of
the caudal peduncle; however, these are less apt
to betray their deception than an erect 1st dorsal
fin.
Sazima (2002b) reported the juvenile of the
snook Centropomus mexicanus to be an aggressive mimic of the mojarra Eucinostomus
melanopterus from observations in southeastern
Brazil. By resembling the mojarra in shape and
317
Fig. 91. Canthigaster margaritata, Red Sea.
color pattern, especially the black-tipped spinous
dorsal fin, the snook is able to closely approach
otherwise wary prey. It often joins foraging mojarras in order to take advantage of prey dislodged
from the substrate by the feeding mojarras. Within
an aggregation of mojarras, a snook may also
enjoy the protection from predators by schooling.
Sazima also found Centropomus parallelus in
aggregations of the mojarras Eucinostomus
melanopterus and E. argenteus and suggested
that this snook may also be an aggressive mimic.
He added that Centropomus unionensis and
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Zoological Studies 44(3): 299-328 (2005)
Fig. 92. Pseudochromis fuscus, Flores, Indonesia.
Fig. 93. Pomacentrus chrysurus, Sumatra, Indonesia.
Fig. 94. Pseudochromis fuscus, Bali, Indonesia.
Fig. 95. Pomacentrus moluccensis, Gulf of Thailand.
Eucinostomus currani of the Pacific coast of South
America may be a mimetic pair as well, judging
from their similarities in color pattern and distribution. He provided in tabular form a review of 39
examples of aggressive mimicry in fishes. He
divided aggressive mimicry into 3 categories:“(1)
fish species that feed on smaller prey than themselves tend to mimic and join fish species harmless to their prospective prey; (2) fish species that
feed on larger prey than themselves tend to mimic
mostly beneficial fish species (cleaners) or, less
frequently, join species harmless to their prospective prey; (3) fish species that feed on prey about
their own size tend to mimic their prospective prey
,
species, the“wolf in a sheep s clothes”disguise
type.”
,
One of Sazima s 39 examples is from the 3rd
category, the carangid Oligoplites saurus that mimics the atherinid Atherinella brasiliensis (Sazima
and Uieda 1980). The carangid joins aggregations
of its model and feeds by removing scales from its
prey. This is the only known marine example of
his type 3 aggressive mimicry.
One would not think that the dottyback
Pseudochromis fuscus could be an aggressive
mimic of damselfishes of the genus Pomacentrus,
but Munday et al. (2003) demonstrated this from
their field research on the Great Barrier Reef.
Pseudochromis fuscus occurs in 2 distinctly different color morphs: brown (Fig. 92), which is colored
like Pomacentrus chrysurus (Fig. 93); and yellow
(Fig. 94), which most resembles either P.
amboinensis or P. moluccensis (Fig. 95). They
noted in the field that the brown dottyback associates with P. chrysurus, and the yellow one with the
yellow damselfishes. Multiple-choice experiments
in the laboratory revealed that the dottyback preferentially selects habitat patches occupied by
damselfishes of the same coloration. Pseudochromis fuscus is a predator of small fishes,
including newly recruited damselfishes.
The example by McCosker (1977), who
depicted the fright posture of the Indo-Pacific plesiopid fish Calloplesiops altivelis, commonly known
as the Comet, is unique. It hides in the reef by day
and emerges at dusk. When threatened, it moves
headfirst into a hole but leaves the posterior part of
its body exposed and expands its fins. It then
resembles the head of the moray eel Gymnothorax
meleagris, with an ocellated black spot at the rear
,
base of the dorsal fin which looks like the moray s
eye. This might seem disputable as an example of
Randall -- Mimicry in Marine Fishes
319
Fig. 96. Novaculoides macrolepidotus, subadult, Kenya
(Andrew Spreinat).
Fig. 97. Ablabys taenianotus, Mindoro, the Philippines
(Andrew Spreinat).
mimicry until one witnesses the behavior.
Photographs of the mimic and model made the
cover of Science (vol. 97).
Randall and Spreinat (2004) reported the
subadult of the Seagrass Wrasse Novaculoides
macrolepidotus (Fig. 96) to be a mimic of venomous waspfishes of the genus Ablabys. Their
illustration of the model is the Cockatoo Waspfish
(A. taenianotus) (Fig. 97). The wrasse (adult
shown above as Fig. 14) is usually very cryptic, but
the individual of figure 96 was exposed and very
sedentary, rocking to and fro like a species of
Ablabys, and holding its dorsal fin fully erect. It
was easily photographed at close range because
of its reluctance to move. The fish is also an
example of protective coloration in that it resembles a fragment of plant material as well as a
waspfish.
Angelfishes of the genus Centropyge are
mimicked by young of the surgeonfish Acanthurus
pyroferus (Fig. 98). When first observed and collected in Tahiti in 1956, the author thought it was a
Fig. 98. Acanthurus pyroferus, juvenile, Line Is.
Fig. 99. Centropyge flavissima, Society Is.
Fig. 100. Acanthurus pyroferus, large juvenile, Tahiti.
Fig. 101. Acanthurus pyroferus, adult, Marshall Is.
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Zoological Studies 44(3): 299-328 (2005)
Fig. 102. Acanthurus pyroferus, juvenile, Scott Reef, Western
Australia.
Fig. 103. Centropyge vrolikii, Ambon, Indonesia.
Fig. 104. Acanthurus tristis, Bali, Indonesia.
Fig. 105. Acanthurus tristis, juvenile, Sumatra.
new species of acanthurid. Its striking resemblance to the common small angelfish Centropyge
flavissima (Fig. 99) immediately suggested mimicry, but it was not clear which was the mimic and
which was the model until individuals of the surgeonfish were seen that were beginning to turn
brown (Fig. 100), and eventually became the typical A. pyroferus adult (Fig. 101). Knowing that
juvenile angelfishes of some species of the genera
Holacanthus and Pomacanthus opportunistically
clean reef fishes, the little C. flavissima was suspected of being a part-time cleaner and therefore
protected from predation. Underwater observation
and analysis of gut contents of 50 specimens soon
disproved that, as no ectoparasites were found.
Most of the food material was algal, with some
sponge. Then the possibility that the angelfish
might have some quality which repels predators
was tested, but the fish were readily eaten when
speared specimens were offered to jacks and
groupers. In writing of the results of that study,
Randall and Randall (1960) admitted to not knowing the biological basis of the mimicry. Years later,
after experiencing the difficulty of taking underwater photographs of species of Centropyge, the
author determined that these small angelfishes are
Fig. 106. Centropyge eibli, Bali, Indonesia.
extremely wary and quickly take cover in the reef
when approached. Predators presumably learn
the same and target easier prey.
In the more western part of the Pacific, the
young of Acanthurus pyroferus (Fig. 102) usually
mimic Centropyge vrolikii (Fig. 103), the most common species of the genus in Australia and New
Guinea (Allen et al. 1998), ranging north to Japan
and west to Sumatra. Acanthurus tristis (Fig. 104),
a close relative of A. pyroferus, occurs in the eastern Indian Ocean, including the Andaman Sea and
islands of southwestern Indonesia. There the
Randall -- Mimicry in Marine Fishes
juvenile (Fig. 105) mimics Centropyge eibli (Fig.
106).
Another explanation for the benefit of the
mimicry of Centropyge vrolikii by the young of
Acanthurus pyroferus was offered by Eagle and
Jones (2004) from observations of these 2 species
and the territorial damselfish Plectroglyphidodon
lacrymatus in Papua New Guinea. They found
that the mimic surgeonfish occurs within 1~2 m of
similar-sized individuals of the angelfish. The surgeonfish was attacked less frequently by the
aggressive damselfish than were other surgeonfishes. The most abundant food category of the
angelfish was reported to be sponge material,
whereas the diet of the mimic surgeonfish was predominately detrital material and sediment, with
small amounts of algae. This was more similar to
the food of P. lacrymatus, so Eagle and Jones
hypothesized,“Mimics may be deceiving the damselfish as to their true diet, which more closely
matches the diet of the damselfish than the diet of
the angelfish.” Sponges dominate the food habits
of angelfishes of the genera Holacanthus and
Pomacanthus (Randall and Hartman 1968), but
reports of the food habits of species of Centropyge
have not shown that sponges are an important
food source. Hiatt and Strasburg (1960) examined
the gut contents of 4 C. flavissima from the
Marshall Is. and reported it to be entirely herbivorous. Randall (1967) found only algae and detritus
in 5 specimens of C. argi from the West Indies.
Hobson (1974) examined the gut contents of 5
specimens of the endemic Hawaiian C. potteri.
Filamentous algae were the major food source, but
there was much unidentified debris, including sand
and foraminiferans; sponges were found in small
amounts (with a ranking index of 2.3, compared to
41.7 for algae and 42.3 for the unidentified debris).
I examined the stomach contents of 3 specimens
of C. potteri after collecting them in 1971 and
1972. I found mainly filamentous algae, some
detritus, a little fine sediment, and no sponge
material. I examined the stomach contents of 8
Bishop Museum specimens of C. vrolikii for this
report. Only 1 from Ishigaki, Ryukyu Is. had more
than 50% of the food composition as sponge material. The others from the Great Barrier Reef,
Indonesia, Palau, and Okinawa had eaten mainly
algae and detritus (probably mostly diatoms); the
stomachs of only 3 of these specimens contained
a trace of sponge. It is suggested that the sample
size for the food-habit study by Eagle and Jones
(2004) was small, and unusual in containing so
much sponge material. The reason for the fewer
321
Fig. 107. Acanthurus pyroferus, juvenile, Vanuatu (Larry
Sharron).
Fig. 108. Centropyge flavissima x C. vrolickii, Marshall Is.
attacks by the damselfish on juvenile Acanthurus
pyroferus is more likely a benefit of less aggression towards the model angelfish because of the
wary behavior of the latter, which darts to the shelter of the reef when threatened. Food habits, however, are an important consideration. Becsuse the
surgeonfish is more restricted in its diet to filamentous algae, it probably requires a broader feeding
area than the angelfish and is therefore more
exposed to predation.
There is a question as to whether Acanthurus
pyroferus, which mimics Centropyge flavissima in
the central and eastern islands of the South
Pacific, is the same species as the one in the
western Pacific where its young usually resemble
C. vrolikii. In an area where both species of
Centropyge occur, and there is only 1 adult form of
Acanthurus pyroferus, it would seem that the juvenile surgeonfish has the capability of mimicking
either species of Centropyge, perhaps depending
on which one is most common in the area of its
settlement on the reef. Evidence for this came
from Vanuatu where both species of Centropyge
occur, as well as hybrids of the two. Juveniles of
Acanthurus pyroferus have been collected by Larry
Sharron at Vanuatu that are mimics (Fig. 107) of
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Zoological Studies 44(3): 299-328 (2005)
Fig. 109. Labroides dimidiatus, Papua New Guinea (Gerald R.
Allen).
Fig. 110. Aspidontus taeniatus, Alor, Indonesia.
the hybrid (Fig. 108).
Barnard (1927) was the first to report the
amazing resemblance of the wrasse Labroides
dimidiatus to the fangblenny Aspidontus taeniatus
from observations in South Africa. He noted that
there are 2 varieties of each species, one with a
dark band across the base of the pectoral fin and
one without. He wrote,“It would be premature to
claim this as a case of mimicry until observations
have been made on the habits of these two
species and on the possible poisonous qualities of
one or the other.”
Randall (1955) also observed the close similarity of Labroides dimidiatus (Fig. 109) and
Aspidontus taeniatus (Fig. 110) from fieldwork in
the Gilbert Is. (Kiribati) in 1951. He described the
unusual mode of swimming of the wrasse, which
oscillates the posterior part of its body up and
down as if to attract attention, and he saw it picking at the bodies of other fishes. He found
calagoid copepods and small isopods in the stomachs of the wrasse and suggested that the blenny
is mimicking it because of protection from predation gained by the wrasse due to its parasite-feeding habit.
As a result of field research in Moorea,
Society Is. in 1956, Randall and Randall (1960)
reported that Aspidontus taeniatus feeds in part on
the fins of reef fishes. In its guise of the Cleaner
Wrasse, and by remaining in the vicinity of a cleaning station, it is able to more closely approach its
prey (often juvenile or subadult reef fishes, rather
than adults that have learned to avoid it).
Juveniles of Labroides dimidiatus are black except
for a bright blue dorsal stripe, and they gradually
change to the adult pattern with growth. The
Mimic Blenny matches the color the Cleaner
Wrasse of the same size as it grows.
Fig. 112. Labroides dimidiatus, Fiji (Gerald R. Allen).
Fig. 113. Aspidontus taeniatus, Fiji (Gerald R. Allen).
Fig. 111. Labroides dimidiatus, Tuamotu Archipelago.
Randall -- Mimicry in Marine Fishes
There is a common 2nd color form of
Labroides dimidiatus in the Society Is. and the
Tuamotu Archipelago that has a salmon pink area
near the middle of the body beneath the black
stripe (Fig. 111). Aspidontus taeniatus lurking in
the vicinity of a Cleaner Wrasse of this color pattern on the fringing reef of Moorea was observed
to exhibit the same color variation. Does the
Mimic Blenny adopt a permanent color pattern to
match the Labroides model in the area where it is
resident, or is it able to alter its pattern if it finds
itself in the area with a model of a different color?
Catching the Mimic Blenny and switching it to an
area with a normal-colored individual, and vice
versa, was considered to see if the 2 blennies
would change their color to match the new models
of Labroides. This was not attempted because
there was too much opportunity for the displaced
blennies to relocate along the linear reef. Ideally,
such a transplantation should be made in a lagoon
with well-isolated patch reefs, or in large aquaria.
The author collected a specimen of a blenny
of the genus Aspidontus at Ua Pou in the
Marquesas Is. in 1971 that was whitish, becoming
yellow posteriorly and on the caudal fin; its dark
stripe was not as black as that of A. taeniatus, and
it had a series of white dots along the upper margin. Typical Aspidontus taeniatus and its usual
model, Labroides dimidiatus, are present in the
Marquesas. Because the specimen seemed to
represent a new species, it was sent to William F.
Smith-Vaniz. He identified it as A. taeniatus and
illustrated it in an update of his review of sabertooth blennies (Smith-Vaniz 1987: fig. 1C). It is
now realized that the fish that served as the
,
blenny s model was the female of the wrasse Coris
hewetti, at that time undescribed (see Randall
1999: pl. 20, fig. C).
A color variant of Labroides dimidiatus with a
posterior section of the black stripe replaced by
bright yellow (Fig. 112) occurs in the South Pacific
from the Samoa Is. to the Coral Sea. No morphological differences could be found to separate it
from typically colored specimens of L. dimidiatus
with which it coexists; however a DNA analysis is
suggested to ascertain whether these are merely
color variants. Individuals of Aspidontus taeniatus
near the yellow-marked Cleaner Wrasse have the
same color pattern (Fig. 113).
A short-term change in color pattern was
demonstrated for the juvenile of the fangblenny
Plagiotremus rhinorhinchos. Like Aspidontus taeniatus, this species is regarded as a mimic of
Labroides dimidiatus, although as an adult (Fig.
323
Fig. 114. Plagiotremus rhinorhynchos, Wetar, Indonesia.
Fig. 115. Plagiotremus rhinorhynchos, juvenile, Flores,
Indonesia.
Fig. 116. Labroides dimidiatus, juvenile, Moyo I., Indonesia.
114), it is not as precise a mimic as A. taeniatus.
Nevertheless, it enjoys a higher striking rate in its
attempts to feed on the skin of reef fishes when in
the proximity of a Cleaner Wrasse (Côte and
Cheney 2004). The juvenile of A. taeniatus (Fig.
115) more closely matches the color of juvenile L.
dimidiatus (Fig. 113) (Kuwamura 1981). In an
impressive field experiment in Kimbe Bay, New
Britain, Moland and Jones (2004) found 92% of
mimetic P. rhinorhinchos closely associated with
juvenile L. dimidiatus. When the Cleaner Wrasse
324
Zoological Studies 44(3): 299-328 (2005)
Fig. 117. Ecsenius midas, Flores, Indonesia.
Fig. 118. Pseudanthias squamipinnis, Red Sea.
Fig. 119. Pseudanthias bartlettorum, Line Is.
Fig. 120. Pseudanthias dispar (above) and Luzonichthys
whitleyi, Line Is.
Fig. 121. Lepidozygus tapeinosoma, Line Is.
Fig. 122. Ecsenius midas, Line Is.
was present, 80% of the fangblenny attacks on
prey fishes were successful. When they removed
the Cleaner Wrasse, the fangblenny began to
change color the 1st day, and by the 7th day it had
reverted to its non-mimetic pattern. The success
rate of its attacks dropped by 10% the 1st day, and
to 20% by the 7th day.
Plagiotremus rhinorhinchos relies heavily on
removing mucus and epidermal tissue of prey fishes for its food. Aspidontus taeniatus feeds much
less frequently by biting pieces from the fins of
other fishes, in spite of its closer resemblance to
Labroides dimidiatus in color, morphology, and
mode of swimming. It also feeds on demersal
eggs of fishes, and sabellid worm tentacles
(Randall and Randall 1960, Kuwamura 1983).
Dafni and Diamant (1984) proposed the term
school-oriented mimicry when a normally solitary
fish species mingles with a similar aggregating
species for the advantage of schooling. Their
example was the juvenile of the fangblenny
Meiacanthus nigrolineatus, which is colored like
dark-striped apogonid fishes with a basal caudal
spot, and joins their aggregations. Other examples are the blenny Ecsenius midas (Fig. 116)
which schools with Pseudanthias (Fig. 117) (Starck
Randall -- Mimicry in Marine Fishes
325
Fig. 123. Pseudanthias tuka and Luzonichthys waitei,
Sulawesi.
Fig. 124. Lutjanus kasmira and Mulloidichthys mimicus,
Marquesas Is. (John L. Earle).
1969), and the apogonid Ostorhinchus compressus, which closely resembles juvenile cardinalfish
of the species Cheilodipterus quinquelineatus and
may aggregate with it (Randall et al. 1990, Kuiter
1991, Allen et al. 2003).
Randall and McCosker (1993) adopted the
term social mimicry for fishes, to replace schooloriented mimicry, because the former is in use for
mixed flocks of birds. Their example in fishes consisted of the 3 anthiine fishes Pseudanthias
bartlettorum (Fig. 118), P. dispar (Fig. 119), and
Luzonichthys whitleyi (Fig. 120), the damselfish
Lepidozygus tapeinosoma (Fig. 121), and the
blenniid Ecsenius midas (Fig. 122), all with a common yellow and pink color pattern, and which
occur in variously mixed schools in the Line Is. and
Phoenix Is. These species feed on zooplankton in
aggregations above the substratum. The bicolored
pattern is the normal coloration only for P. bartlettorum and L. whitleyi. Randall and McCosker also
illustrated Pseudanthias tuka and Luzonichthys
waitei in a mixed aggregation (Fig. 123) as social
mimics (in their female color pattern).
Randall and Guézé (1980) described the
goatfish Mulloidichthys mimicus from the
Marquesas Is. and Line Is.; its species name is in
reference to its close resemblance to the snapper
Lutjanus kasmira with which it often schools (Fig.
124). The snapper is mainly nocturnal and tends
to form semi-stationary aggregations by day.
Krajewski et al. (2004) presented evidence,
including color photos, to show that the western
Atlantic Yellow Goatfish Mulloidichthys martinicus
may be seen in schools with the Smallmouth Grunt
Haemulon chrysargerum, and is therefore a similar
example of social mimicry. One of their photographs shows a previously inactive aggregation
of the goatfish joining a school of the grunt when
frightened by an approaching diver.
Randall (1998) wrote that herbivorous surgeonfishes of the genus Acanthurus may form
feeding aggregations of more than 1 species in
order to overcome territorial damselfishes trying to
protect their private pastures of benthic algae. He
provided an illustration of a mixed school of A.
coeruleus and A. chirurgus off the Caribbean coast
of Honduras.
Another example of social mimicry was published by Bunkley-Williams and Williams (2000).
They observed the juvenile of the Black Snapper
Apsilus dentatus swimming with a feeding aggregation of Blue Chromis Chromis cyanea in 12~27 m
on the outer-reef slope in the Cayman Is. Unlike
the adult snapper, the juveniles were colored like
the damselfish, i.e., blue with a broad black upper
and lower margin of the caudal fin. Although
slightly larger than the chromis, the snapper were
difficult to distinguish. Both the chromis and juvenile snapper feed on zooplankton.
Smith-Vaniz et al. (2001) regarded the fangblenny Meiacanthus urostigma as a social mimic of
the cardinalfish Cheilodipterus quinquelineatus.
Both have black stripes and a broad yellow area
around a black spot near the base of the caudal
fin. The 2nd author of that study, Ukkrit
Satapoomin, observed young M. urostigma aggregating with juveniles of C. quinquelineatus.
Acknowledgments: I am very grateful to Gerald
R. Allen for supplying 7 photographs for this report,
John L. Earle, Richard Field, Paul Humann, and
Andreas Spreinat for 2 photographs, and to the following for 1 photo each (credit line with the photographs): R. Charles Anderson, David Hall, Yves
Lefevre (via Philippe Bacchet), Hajime Masuda,
Robert F. Myers, Leslie Newman, Gustav Paulay,
326
Zoological Studies 44(3): 299-328 (2005)
Barry C. Russell, Frank Schneidewind, Mike
Severns, Larry Sharron, Victor G. Springer, and
Ron and Valerie Taylor. Dannie A. Hensley, Ivan
Sazima, and William F. Smith-Vaniz provided overlooked references. The manuscript was reviewed
by Gerald R. Allen, John L. Earle, Richard L. Pyle,
Helen A. Randall, Ivan Sazima, William F. SmithVaniz, and Victor G. Springer, all of whom made
helpful comments.
REFERENCES
Allen GR, BC Russell, BA Carlson, WA Starck II. 1975.
Mimicry in marine fishes. Trop. Fish Hobbyist 25: 47-56.
Allen GR, R Steene, M Allen. 1998. A guide to angelfishes
and butterflyfishes. Odyssey Publishing; Australia:
Tropical Reef Research.
Allen GR, R Steene, P Humann, N DeLoach. 2003. Reef fish
identification tropical Pacific. Jacksonville, FL: New World
Publications; El Cajon, CA: Odyssey Publishing.
Amaoka K, H Senou, A Ono. 1994. Record of the bothid flounder Asterorhombus fijiensis from the western Pacific, with
observations on the use of the first dorsal-fin ray as a lure.
Jpn. J. Ichthyol. 41: 23-28.
Barnard KH. 1927. A monograph of the marine fishes of South
Africa. Ann. S. Afr. Mus. 21: v + 419-1065.
Breder CM Jr. 1946. An analysis of the deceptive resemblances of fishes to plant parts, with critical remarks on
protective coloration, mimicry and adaptation. Bull.
Bingham Oceanogr. Coll. 10: 1-49.
Bunkley-Williams L, EH Williams Jr. 2000. Juvenile Black
Snapper, Aspilus dentatus (Lutjanidae), mimic Blue
Chromis, Chromis cyanea (Pomacentridae). Copeia
2000: 579-581.
Caley MJ, D Schluter. 2003. Predators favour mimicry in a
tropical reef fish. Proc. Roy. Soc. Lond. B 270: 667-672.
Clark E, HAF Gohar. 1953. The fishes of the Red Sea: Order
Plectognathi. Publ. Mar. Biol. Stat. Al Ghardaqa Egypt 8:
1-80.
Côte IM, KL Cheney. 2004. Distance-dependent costs and
benefits of aggressive mimicry in a cleaning symbiosis.
Proc. Biol. Sci. 271: 2627-2630.
Cott HB. 1957. Adaptive coloration in animals. London:
Methuen and Co.
Cuvier G, A Valenciennes. 1829. Histoire Naturelle des
Poissons. Vol. 3. Paris: F.G. Levrault.
Dafni J, A Diamant. 1984. School-oriented mimicry, a new
type of mimicry in fishes. Mar. Ecol. Prog. Ser. 20: 45-50.
Domeier ML. 1994. Speciation in the serranid fish genus
Hypoplectrus. Bull. Mar. Sci. 54: 103-141.
Eagle JV, GP Jones. 2004. Mimicry in coral reef fishes: ecological and behavioural responses of a mimic to its model.
J. Zool. Lond. 264: 33-43.
Field R. 1997. Mimicry in Red Sea reef fishes. J. Saudi Arab.
Nat. Hist. Soc. 3: 32-35.
Fisher RA. 1930. The genetical theory of natural selection.
Oxford, UK: Clarendon Press.
Goren M, I Karplus. 1983. Preliminary observations on the
scorpion fish Scorpaenodes guamensis and its possible
mimic the cardinal fish Fowleria abocellata. Develop.
Ecol. Environ. Qual. 2: 327-336.
Heck KL Jr, MP Weinstein. 1978. Mimetic relationships
between tropical burrfishes and opisthobranchs.
Biotropica 10: 78-79.
Hiatt RW, DW Strasburg. 1960. Ecological relationships of the
fish fauna on coral reefs of the Marshall Islands. Ecol.
Monogr. 30: 65-127.
Hobson ES. 1969. Possible advantages to the blenny Runula
azalea in aggregating with the wrasse Thalassoma
lucasanam in the tropical eastern Pacific. Copeia 1969:
191-193.
Hobson ES. 1974. Feeding relationships of teleostean fishes
on coral reefs in Kona, Hawaii. Fish. Bull. 72: 915-1031.
Krajewski JP, RM Bonaldo, C Sazima, I Sazima. 2004. The
association of the goatfish Mulloidichthys martinicus with
the grunt Haemulon chrysargyreum: an example of protective mimicry. Biota Neotrop. 4: 1-4.
,
Kuiter RH. 1991. Nature s copies. Sportdiving no. 23: 115116.
Kuiter RH. 1995. The juvenile vermicular cod Plectropomus
oligacanthus, a mimic of the slender Maori wrasse
Cheilinus celebicus. Rev. Fr. Aquariol. 21: 3-4.
Kuwamura T. 1981. Mimicry of the cleaner wrasse Labroides
dimidiatus by the blennies Aspidontus taeniatus and
Plagiotremus rhinorhynchos. Nankiseibutu Nanki Biol.
Soc. 23: 61-70. (in Japanese with English summary)
Kuwamura T. 1983. Reexamination on the aggressive mimicry
of the cleaner wrasse Labroides dimidiatus by the blenny
Aspidontus taeniatus (Pisces: Perciformes). J. Ethol. 1:
22-33.
Longley WH. 1917. Studies upon the biological significance of
animal coloration II. A revised working hypothesis of mimicry. Am. Nat. 51: 257-285.
Longley WH, SF Hildebrand. 1940. New genera and species
of fishes from Tortugas, Florida. Pap. Tortugas Lab.
(Carnegie Inst. Wash.) 32: 225-285.
Losey GS. 1972. Predator protection in the poison fang
blenny, Meiacanthus atrodorsalis, and its mimics,
Ecsenius bicolor and Runula laudandus (Blenniidae).
Pac. Sci. 26: 129-139.
Mahadevan S, K Nagappan Nayaar. 1965. Underwater ecological observations in the Gulf of Mannar, off Tuticorin. J.
Mar. Biol. Assoc. India 7: 197-199.
Manabe H, A Shinomiya. 1998. Use of the first dorsal fin ray
as a lure by the bothid flounder, Aserorhombus intermedius. Jpn. J. Ecol. 48: 117-121 (in Japanese)
Mansueti R. 1963. Symbiotic behavior between small fishes
and jellyfishes, with new data on that between the stromateid, Peprilus alepidotus, and the scyphomedusa,
Chrysaora quinquecirrha. Copeia 1963: 40-80.
Masterman AT. 1908. On a possible case of mimicry in the
common sole. J. Linn. Soc. (Zool.) 30: 239-244.
McCosker JE. 1977. Fright posture of the plesiopid fish
Calloplesiops altivelis: an example of Batesian mimicry.
Science 197: 400-401.
McCosker JE. 1982. A new genus and two new species of
remarkable Pacific worm eels (Ophichthidae, subfamily
Myrophinae). Proc. CA Acad. Sci. 43: 59-66.
McCosker JE, RH Rosenblatt. 1993. A revision of the snake
eel genus Myrichthys (Anguilliformes: Ophichthidae) with
the description of a new eastern Pacific species. Proc.
CA Acad. Sci. 48: 153-169.
Moland E, GP Jones. 2004. Experimental confirmation of
aggressive mimicry by a coral reef fish. Oecologia 140:
676-683.
Moyer JT. 1977. Aggressive mimicry between juveniles of the
snapper Lutjanus bohar and species of the damselfish
Randall -- Mimicry in Marine Fishes
genus Chromis from Japan. Jpn. J. Ichthyol. 24: 218-222.
Munday PL, PJ Eyre, GP Jones. 2003. Ecological mechanisms for coexistence of colour polymorphism in a coralreef fish: an experimental evaluation. Oecologia 442:
519-526.
Newman L, L Cannon. 2003. Marine flatworms The world of
polyclads. Collingswood, Victoria, Australia: CSIRO
Publishing.
Ormand RFG. 1980. Aggressive mimicry and other interspecific feeding associations among Red Sea coral reef
predators. J. Zool. Lond. 191: 247-262.
Pietsch TW, DB Grobecker. 1978. The angler: aggressive
mimicry in an antennariid anglerfish. Science 201: 369370.
Pietsch TW, DB Grobecker. 1987. Frogfishes of the world.
Stanford, CA: Stanford Univ. Press.
Poulton WB. 1898. Natural selection the cause of mimetic
resemblance and common warning colors. J. Linn. Soc.
(Zool.) 26: 558-612.
Quoy JRC, P Gaimard. 1824-1826. Voyage autour du Monde
,
- Exécuté sur les Corvettes de S.M.“l Uranie”et“La
Physicienne”pendant les Années 1817-1820. Paris:
Pillet Aîné, pp. 192-401.
Randall JE. 1955. Fishes of the Gilbert Islands. Atoll Res.
Bull. 47: xi + 243.
Randall JE. 1967. Food habits of reef fishes of the West
Indies. Stud. Trop. Oceanogr. no. 5: 665-847.
Randall JE. 1995. Coastal fishes of Oman. Honolulu, HI: Univ.
,
of Hawai i Press.
Randall JE. 1998. Caribbean reef fishes, ed. 3. Neptune City,
NJ: T.F.H. Publishing.
Randall JE. 1999. Revision of the Indo-Pacific labrid fishes of
the genus Coris, with descriptions of five new species.
Indo-Pac. Fishes 29: 1-74.
Randall JE. 2002. Surgeonfishes of the world. Honolulu, HI:
Mutual Publishing and Bishop Museum Press.
Randall JE. 2005. Reef and shore fishes of the South Pacific.
,
Honolulu, HI: Univ. of Hawai i Press.
Randall JE, GR Allen, RC Steene. 1990. Fishes of the Great
Barrier Reef and Coral Sea. Honolulu, HI: Univ. of
Hawai`i Press.
Randall JE, JL Earle. 2004. Novaculoides, a new genus for
the Indo-Pacific labrid fish Novaculichthys
macrolepidotus. Aqua, J. of Ichtoyol. and Aquatic Biol. 8:
37-43.
Randall JE, AR Emery. 1971. On the resemblance of the
young of the fishes Platax pinnatus and Plectorhynchus
chaetodontoides to flatworms and nudibranchs.
Zoologica 56: 115-119.
Randall JE, P Guézé. 1980. The goatfish Mulloidicthys mimicus, n. sp. (Pisces: Mullidae) from Oceania, a mimic of
the snapper Lutjanus kasmira (Pisces: Lutjanidae). Bull.
Mus. Natl. Hist. Nat. (Paris), 4 sér., 2: 603-609.
Randall JE, WD Hartman. 1968. Sponge-feeding fishes of the
West Indies. Mar. Biol. 1: 216-225.
Randall JE, RH Kuiter. 1989. The juvenile Indo-Pacific grouper
Anyperodon leucogrammicus, a mimic of the wrasse
Halichoeres purpurescens and allied species, with a
review of the recent literature on mimicry in fishes. Rev.
Fr. Aquariol. 16: 51-56.
Randall JE, JE McCosker. 1993. Social mimicry in fishes.
Rev. Fr. Aquariol. 20: 5-8.
Randall JE, HA Randall. 1960. Examples of mimicry and protective resemblance in tropical marine fishes. Bull. Mar.
Sci. Gulf Carib. 10: 444-480.
327
Randall JE, A Spreinat. 2004. The subadult of the labrid fish
Novaculoides macrolepidotus, a mimic of waspfishes of
the genus Ablabys. Aqua 8: 45-48.
Roberts TR. 1990. Mimicry of prey by fin-eating fishes of the
African characoid genus Eugnathichthys (Pisces:
Distichodidae). Ichthyol. Explor. Freshw. 1: 23-31.
Rocha LA. 2004. Mitochondrial DNA and color pattern variation in three western Atlantic Halichoeres (Labridae), with
revalidation of two species. Copeia 2004: 770-782.
Russell BC. 1975. Disguise, defense and aggression. Austral.
Nat. Hist. 18: 324-329.
Russell BC. 1977. Sabre-teeth and poison-fanged blennies.
Oceans 1: 66-69.
Russell BC, GR Allen, HR Lubbock. 1976. New cases of mimicry in marine fishes. J. Zool. (Lond.) 180: 407-423.
Sazima I. 2002a. Juvenile grunt (Haemulidae) mimicking a
venomous leatherjacket (Carangidae), with a summary of
Batesian mimicry in marine fishes. Aqua, J. of Ichtoyol,
and Aquatic Biol. 6: 61-68.
Sazima I. 2002b. Juvenile snooks (Centropomidae) as mimics
of mojarras (Gerreidae), with a review of aggressive mimicry in fishes. Environ. Biol. Fishes 65: 37-45.
Sazima I, VS Uieda. 1980. Scale-eating behavior in
Oligoplites saurus and record of scale-eating in O.
palometa and O. saliens (Pisces, Carangidae). Rev.
Bras. Biol. 40: 701-710. (in Portuguese)
Schneidewind F. 2004. Neue Anglerfisch - Mimikry. Aquar.
Fachmag. 35: 86-88.
Seigel JA, TA Adamson. 1983. Batesian mimicry between a
cardinalfish (Apogonidae) and a venomous scorpionfish
(Scorpaenidae) from the Philippine Islands. Pac. Sci. 37:
75-79.
Shallenberger RJ, WD Madden. 1973. Luring behavior in the
scorpionfish, Iracundus signifer. Behaviour 47: 3-47.
Sikkel PC, PD Hardison. 1992. Interspecific feeding associations between the goatfish Mulloides martinicus (Mullidae)
and a possible aggressive mimic, the snapper Ocyurus
chrysurus (Lutjanidae). Copeia 1992: 914-917.
Smith-Vaniz WF. 1976. The saber-toothed blennies, tribe
Nemophini (Pisces: Blenniidae). Monogr. Acad. Nat. Sci.
Phil. 19: 1-196.
Smith-Vaniz WF. 1987. The saber-toothed blennies, tribe
Nemophini (Pisces: B lenniidae): an update. Proc. Acad.
Nat. Sci. Phil. 139: 1-52.
Smith-Vaniz WF, U Satapoomin, GR Allen. 2001. Meiacanthus
urostigma, a new fangblenny from the northeastern Indian
Ocean, with discussion and examples of mimicry in
species of Meiacanthus (Teleostei: Blenniidae:
Nemophini). Aqua, J. of Ichthyol. and Aquatic Biol. 5: 2543.
Snyder DB. 1999. Mimicry of initial-phase bluehead wrasse,
Thalassoma bifasciatum (Labridae) by juvenile tiger
grouper, Mycteroperca tigris (Serranidae). Rev. Fr.
Aquariol. 26: 17-20.
Snyder DB, JE Randall, SW Michael. 2001. Aggressive mimicry by the juvenile of the redmouth grouper, Aethaloperca
rogaa. Cybium 25: 227-232.
Springer VG. 1988. The Indo-Pacific blenniid fish genus
Ecsenius. Smithson. Contr. Zool. 465: 1-134.
Springer VG, WF Smith-Vaniz. 1972. Mimetic relationships
involving fishes of the family Blenniidae. Smithson. Contr.
Zool. 112: 1-36.
Starck WA II. 1969. Ecsenius (Anthiiblennius) midas, a new
subgenus and species of mimic blenny from the western
Indian Ocean. Not. Nat. (Phil.) 419: 1-9.
328
Zoological Studies 44(3): 299-328 (2005)
Thresher RE. 1978. Polymorphism, mimicry, and the evolution
of the hamlets (Hypoplectrus, Serranidae). Bull. Mar. Sci.
28: 345-353.
Tyler JC. 1966. Mimicry between the plectognath fishes
Canthigaster valentini (Canthigasteridae) and Paraluteres
prionurus (Aluteridae). Not. Nat. (Phil.) 386: 1-13.
Vane-Wright RI. 1980. On the definition of mimicry. Zool. J.
Linn. Soc. 13: 1-6.
Whitley GP. 1935. Fishes from Princess Charlotte Bay, North
Queensland. Rec. S. Austral. Mus. 5: 345-363.
Wickler W. 1965. Mimicry and the evolution of animal communication. Nature 208: 519-521.
Wickler W. 1968. Mimicry in plants and animals. New York,
NY: McGraw-Hill.
Yatsu A, A Iwata, M Sato. 1983. First records of the blenniid
fishes, Petroscirtes springeri and Petroscirtes variabilis,
from Japan. Jpn. J. Ichthyol. 30: 297-300.